Pollen evolution and its taxonomic significance in Cuscuta (dodders ...

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M. Welsh et al. clades of subg. Grammica exhibit little pollen variation. (e.g., clades A, B, C, E, H, L, and N, see Stefanović et al. 2007; Table 2). Nevertheless, even in groups such as clade A (C. californica complex, Costea et al. 2006d), clade B (C. pentagona complex, Costea et al. 2006b), and clade L (C. gracillima complex, Costea et al. 2008a), basic pollen characters such as size, shape, and diameter of puncta/perforations have already been used together with other characters to separate species. Pollen will play an increasingly significant role in the subg. Monogynella and many of the Grammica clades (e.g., D, F, G, I, K, and O, see Stefanović et al. 2007) which exhibit significantly more palynological diversity (Table 2). These infrageneric groups are the least known in the genus, and their future species-level taxonomic revisions will benefit enormously from these additional pollen characters. For example, clade O (subg. Grammica), comprising over 20 species distributed mostly in South America (but also 1 in Africa), is perhaps the most diverse and challenging in Cuscuta (Stefanović et al. 2007). Pollen is equally varied in this group, encompassing practically almost entirely the variation documented in the genus (Table 2). While most species are tricolpate, C. boliviana, C. grandiflora, and C. purpurata are 5-8-colpate. Tectum varies from imperforate in C. purpurata to reticulate in C. parodiniana and C. paitana. Additionally, size and shape can also be used to separate closely related species in this clade. Conclusions Placed in the context of the evolutionary history of pollen in Convolvulaceae (3-colpate 5-6-zonocolpate pantocolpate pantoporate), the pollen of Cuscuta can be considered one step above the primitive because some species in two major lineages (subg. Monogynella and clade O in subg. Grammica) have evolved 5–8 zonocolpate pollen, and because pantocolpate grains, although rare, are present. Reticulate pollen has evolved two times in Cuscuta: in subg. Monogynella and clade O of subg. Grammica. The traditional, qualitative tectum ‘‘types’’ represent arbitrary ranges of variation, which in Cuscuta are better described quantitatively. Overall, the morphology of pollen supports Cuscuta as a sister either to the ‘‘bifid-style’’ clade (Dicranostyloideae) or to one of the members of this clade. Although the pollen characters are insufficient to reconstruct the phylogeny of the genus, pollen morphology is useful for the taxonomy at species level. Acknowledgments The authors warmly thank the curators and directors of AAU, ALTA, ARIZ, ASU, B, BAB, BOL, BRIT, CANB, CAS, CEN, CHR, CHSC, CIIDIR, CIMI, CTES, DAO, ENCB, F, G, GH, H, HUFU, IAC, IBUG, IEB, IND, J, JEPS, LL, LP, LPB, LPS, K, MEL, MERL, MEXU, MICH, MO, NMC, NU, NY, OKLA, OSU, OXF, PACA, PRE, QCNE, QFA, P, PACA, RB, RSA, SAM, S, SD, SGO, SI, SPF, TEX, TRT, UA, UB, UBC, UCR, UCT, UNB, UNM, UPRRP, UPS, US, USAS, VEN, WTU, and XAL for supplying plant material. We also thank Frédérique Guinel and two anonymous reviewers for their comments on an earlier version of the manuscript. Thierry Deroin provided the information about the pollen of Humbertia. This research was supported by Discovery grants from the Natural Sciences and Engineering Research Council (NSERC) of Canada to M. Costea (327013–06) and to S. Stefanović (326439–06). Appendix Vouchers used for scanning electron microscopy. Species (arranged in alphabetical order), country, collector, collector number, herbarium acronym from Index Herbariorum. Cuscuta acuta Engelm. Ecuador, Galapagos Islands, Anderson 1853 (S); Fagerlind & Wibom 3401 (S); Howell 10140 (G). C. acutiloba Engelm.: Bolivia, Mardon 1481 (G); Peru, Pennell 13242 (USM). C. africana Thumb.: South Africa, Beyers 6968 (NBG); Oliver 11852 (NBG). C. americana L.: USA, Florida, Small et al. 11596 (NY); Mexico, Van Devender and Reina-G 2000-745 (WLU); Bahamas, Correll 45030 (NY); Cuba, Britton et al. 5917 (NY). C. angulata Engelm.: South Africa, Beyers 6968/13 (NBG); Orchard 460 (NU). C. appendiculata Engelm.: South Africa, Hofmeyr s.n. (GRA); Burrows 4666 (J); Bohnen 7827 (NBG). C. applanata Engelm. USA, New Mexico, Casteller 7339 (UNM); Mexico, Van Devender 2001-710 et al. (WLU). C. approximata Bab.: USA, California, Abrams 457 (CAS); Mexico, Henrickson 13083a (RSA); Puebla, McKee 11042 (MEXU). C. argentiniana Yunck.: Argentina, Krapovickas & Schinini 36049, 36757 (CTES); Brücher s.n. (S); Meyer 12467 (UPS). C. aurea Liebm.: Mexico, Palmer 87 (S); Guerrero, Duran & Garcia 634 (MEXU). C. australis R. Br. var. australis: New Caledonia, Bonati 737 (S); Australia, Conveny 756 (RSA). C. australis var. tinei (Insenga) Yunck.: Hungary, Simonkai 2635 (NY); Karkovány s.n. (WLU). C. bella Yunck.: Peru, Killip & Smith 21827 (US). C. boldinghii Urb.: Mexico, Van Devender 92-31 et al. (ARIZ); Sinaloa, Vega 2630 (MEXU). C. boliviana Yunck.: Argentina, Hunzinker 2676 (S); Ruiz Leal 14817 (MERL), Burkart 12503 (CTES). C. brachycalyx Yunck.: USA, California, Ahart 9856 (CHICO); Colwell AC 04-305 (YM/WLU). C. burrelli Yunck.: Brazil, Heringer et al. 43 (UB); Alvarenga-Pereira 766 (RB). C. californica Hook. & Arn: USA, California, Sanders 25122 (UCR); Pinzl 7238a (NY); White 5033 (ASU). C. campestris Yunck.: USA, Iowa, Fay 4568 (UC); Oklahoma, Lipscomb 1894 (SMU); Puerto Rico, Liogier & Martorell 13908 (UPRRP); Venezuela, Killip & Tamayo 37010 (GH). C. cassytoides 123
Pollen evolution and its taxonomic significance in Cuscuta Nees: South Africa, Balkwill 6968 (NU); Alexandre 2407 (NBG). C. cephalanthi Engelm.: USA, Illinois, McDonald s.n. (NMS); Missouri, Steyermark 79977(MO); Nebraska, Churchill 4560 (MO). C. chapalana Yunck.: Mexico, Garcia-Ruiz 7942 (CIMI); Jalisco, Carrillo-Reyes & al. 468 (CIMI). C. chilensis Ker Gawl.: Chile, Anderson 84- 189 (S); Buchtien s.n. (S); Valeutey 94 (S). C. chinensis Lam.: Australia, Carter 628 (CANB); China, Wang Wen- Tsai 2378 (RSA). C. choisiana Yunck.: Mexico, Véase 490 (MEXU). C. cockerellii Yunck.: Peru, Vargas 2600 (CUS). C. colombiana Yunck.: Colombia, Haught 4535 (NY); Venezuela, Aristeguieta 4500 (VEN); Tamayo 4432 (VEN). C. compacta Juss.: USA Mississippi, McDaniel 27291 (MO); Maryland, Hill 17349 (MO); New Jersey, Moldenke & Moldenke 25129 (NY). C. corniculata Engelm.: Brazil, Stannard et al. 51861 (G); Colombia, Pennell 1453 (GH). C. coryli Engelm.: USA, Arkansas, Demaree 19603 (CAS); Maryland, Killip 31293 (NY); Michigan, Hanes 548 (NY); Nebraska, Reynolds 2727; Tennessee, Rydberg 8179 (NY). C. corymbosa Ruiz & Pav. var. grandiflora Engelm.: Mexico, Garcia-Ruiz et al. 7572 (CIMI, WLU); Iltis & Guzman 29077 (MEXU); Galván & Galván 4681 (MEXU). C. corymbosa var. stylosa (Choisy) Engelm.: Mexico, Rzedowski 28752 (UCR); Bopp 206 (MEXU); Pringle s.n. (MEXU). C. costaricensis Yunck.: Mexico, Reina-G. et al. 2006-1049 (ARIZ, WLU); Van Devender 98-1789 (ARIZ); Wiens 96-125 et al. (WLU). C. cotijana Costea & I. García: Mexico, García Ruiz 7412 (CIMI,); García Ruiz et al. 7557 (CIMI); Carranza et al. 7316 (IEB). C. cozumeliensis Yunck.: Guatemala, Kellerman 6580 (F); Mexico, Calzade & Nivea 9427 (XAL); Vazquez 176 (MEXU). C. cristata Engelm.: Meyer 10669 (CTES); Argentina, Cabrera 7144 (NY); Hunzinker 4927 (S); Burkart 14000 (SI). C. cuspidata Engelm.: USA, Arkansas, Demaree 15522 (RSA); Texas, Higgins 12480 (NY); Runyon 2828 (SMU). C. decipiens Yunck.: Mexico, Henrickson 6362, 13394, 22781 (RSA). C. deltoidea Yunck.: Mexico, Orcutt 4457 (MEXU); Pringle 5350 (US); García Ruíz 2516 (CIMI). C. dentatasquamata Yunck.: Mexico, Jones s.n. (RSA); USA, Arizona, Lemmon s.n. (UC). C. denticulata Engelm.: USA, Arizona, Peebles & Parker 14793 (NY); California, Thomas 8904 (UC); Nevada, Perish 10299 (CAS); Utah, Choong s.n. (NY). C. desmouliniana Yunck.: Mexico, Spellenberg 4943 (NMS); Van Devender & Reina-G 2002- 23 (WLU); Van Devender 96-360 et al. (WLU). C. draconella Costea & Stefanović: USA, New Mexico, Spellenberg & Mahrt 10497 (NY); Wagner 3395 (UNM); Herman 462 (NY). C. epilinum Weihe: Canada, Quebec, Barabe 16914 (DAO); Sweden, Samuelson 1317 (RSA). C. epithymum (L.) L.: Argentina, Bana 14733 (CTES); USA, New York, Ahles 67695 (SMU); Belgium, Meulebrouck s.n. (WLU). C. erosa Yunck.: USA, Arizona, Jones s.n. (CAS); Mexico, Rebman 4275; Sánchez NF-172 et al. (WLU). C. europea L.: Finland, Alava et al. s.n. (OSU); Sweden, Holmgren 19784 (SD). C. exaltata Engelm.: USA, Texas, Snyder 472 (SMU); Carter 10584 (MO). C. floribunda aff. Kunth: Mexico, Prather & Soule 1221 (TEX). C. flossdorfii Hicken var. pampagrandensis Yunck.: Bolivia, Mendoza & Acebo 919 (MO). C. foetida Kunth var. foetida: Ecuador, Holm-Nielsen & Andrade 18480 (AAU); Holm-Nielsen et al. 5181 (AAU); Peru, Smith 1624 (MO). C. foetida var. pycnantha Yunck: Peru, Plowman et al. 14291 (F). C. friesii Yunck.: Argentina, Krapovickas et al. 21898 (CTES); Saravia Toledo et al. 12993 (CTES); Mulgura 1245 (SI). C. glabrior (Engelm.) Yunck.: USA Texas, Wolff 3270 (NY); Mexico, Marsh 1115 (SMU); Henrickson 13676 (RSA). C. globiflora Engelm.: Argentina, Mulgura et al. 1199 (MO); Bolivia, Plowman & Davis 5196 (GH); Peru, King et al. 247 (USM). C. globulosa Benth: Puerto Rico, Stahl 1064 (S); Urban 855 (S); Liogier & Oquendo 180 (UPRRP). C. glomerata Choisy: USA, Texas, Barkley 13886 (RSA); Wolff 3321 (SMU). C. goyaziana Yunck.: Brazil, Macedo 3731 (NY). C. gracillima Engelm.: Mexico, Pringle 6716 (NML); Koch and Fryxell 82253 (NY); García Ruiz 7334 (CIMI, WLU). C. grandiflora Kunth: Argentina, Schinini et al. 34615 (CTES); Ecuador, Løjtnant et al. 11829 (AAU); Tipaz et al. 1563 (QCNE); Peru, Pennell 13613 (GH). C. gronovii Willd. ex Roem. & Schult. var. gronovii: Canada, Ontario, Catling 5111 (DAO). USA Alabama, Kpeooer et al. s.n. (NY); Georgia, Mellinger s.n. (SMU). C. gronovii var. latiflora Engelm.: USA, Missouri, Brant & Donnell 4813 (MO); Texas, Lundell 11721 (SMU); New York, Ferguson 6091 (NY). C. gymnocarpa Engelm. Galapagos Islands, Fagerlind & Wibom 3658 (S); Werff 2068 (S); Werff 2136 (NY). C. harperi Small: USA, Alabama, Demaree 46295 (NY); Harper s.n. (NY). C. haughtii Yunck.: Ecuador, Asplund 15974 (S); Holm- Nielsen et al. 2308 (NY). C. howelliana Rubtzoff: USA, California, True 7407 (DS); Oswald & Ahart 7978 (JEPS); Reino & Alava 6809 (JEPS). C. hyalina Roth.: India, Pushpauder s.n. (CANB); Namibia, Bosch 25022 (BOL). C. incurvata Prog.: Paraguay, López et al. 243 (CTES); Anisits 2395 (S); Hassler 8170 (S). C. indecora Choisy var. indecora: USA, California, Wolf 4392 (UC); Louisiana, Allen 19239 (BRIT); New Mexico, Spellenberg et al. 3427 (NY). C. indecora var. attenuata (Waterf.) Costea: USA, Oklahoma, Waterfall 17496 (GH); Texas, Fisher 4118 (CAS); Mexico, Palmer 333 (F). C. indecora var. longisepala Yunck.: Argentina, Leal 7964 (NY); USA Texas, Runyon 2819 (NY). C. jalapensis Schltdl.: Mexico, Ton & Lopez 9826 (NY); Sharp 45380 (NY); Waterfall & Wallis 14213 (SMU). C. japonica Choisy: China, Bartholomew et al. 883 (NY); Japan, Furuse 6890 (RSA). C. jepsonii Yunck.: USA, California, Dudley 1774 (DS); 123
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